.SEXUAL INHERITANCE IN THE VIOLET 

C Viola odorata) 



A THESIS 

Presented to the Faculty of the Graduate School 

OF Cornell University for the degree of 

DOCTOR OF PHILOSOPHY 



BY 



ROY DAVID ANTHONY 



Reprinted from Technical Bulletin No. 76 
New York Agricultural Experiment Station, March, 1920 



ASEXUAL INHERITANCE IN THE VIOLET 

( Viola odor at a) 



A THESIS 

Presented to the Faculty of the Graduate School 

OF Cornell University for the degree of 

DOCTOR OF PHILOSOPHY 



BY 



ROY DAVID ANTHONY 

n 



Reprinted from Technical Bulletin No. 76 
New York Agricultural Experiment Station, March, 1920 







^^^^ 



Digitized by the Internet Archive 
in 2010 with funding from 
The Library of Congress 



http://www.archive.org/details/asexualinheritanOOanth 



TECHNICAL BULLETIN No. 76. 



MARCH, 1920. 



rv 









ASEXUAL DTHERITANCE IN THE VIOLET (Viola odorata). 



ROY D. ANTHONY. 




PUBLISHED BY THE STATION. 



BOARD OF CONTROL. 

Governor Alfred E. Smith, Albany. 
Commissioner Charles S. Wilson, Albany. 
Irving Rouse, Rochester. 
Frank M. Bradley, Barkers. 
Charles C. Sackett, Canandaigua. 
Charles R. Mellen, Geneva. 
John B. Mulford, Lodi. 
C. Fred Boshart, Lowville. 
Parker Corning, Albany. 

OFFICERS OF THE BOARD. 

Commissioner Charles S. Wilson, William O'Hanlon, 

President. Secretary and Treasurer. 

STATION STAFF. 



Whitman H. Jordan, Sc.D., LL.D., 

Director. 
fALBERT R. Mann, A.M., 

Agricultural Economist. 
George W. Churchill, 

Agriculturist and Superin- 
tendent of Labor. 
Reginald C. Collison, M.S., 

Agronomist. 
fT. Ltttleton Lton, Ph.D., 

Chemist (Agronomy). 
James E. Mensching, M.S., 

Associate Chemist (Agronomy). 
James D. Harlan, B.S., 

Assistant Agronomist. 
William P. Wheeler, 

First Assistant (Animal Industry). 
Robert S. Breed, Ph.D., 
fWiLLiAM A. Stocking, Jr., M.S.A., 

Bacteriologists. 
Harold J. Conn, Ph.D., 

Associate Bacteriologist. 
John W. Bright, M.S., 
George J. Hucker, M.A., 

Assistant Bacteriologists. 
Fred C. Stewart, M.S., 
tDoNALD Reddick, Ph.D., Botanists. 
Walter O. Gloter, M.A., 

Associate Botanist. 
Mangel T. Munn, M.S., 

Assistant Botanist. 
Lucius L. Van Slkte, Ph.D., Chemist. 
Rudolph J. Anderson, Ph.D., 
fLEONARD A. Maynard, Ph.D., 

Bio-Chemists. 
Arthur W. Clark, B.S., 
Iichard F. Keeler, A.B., 

Associate Chemists. 
Morgan P. Sweeney, A.M., 
Otto McCreary, B.S., 
William F. Walsh, B.S., 

Assistant Chemists. 



Walter L. Kulp, M.S., , 

Harold L. Winston, B.S., 
Millard G. Moore, B.S., 

Assistant Chemists. 
George A. Smith, Dairy Expert. 

James D. Luckett, M.S. A., 

Editor and Librarian. 
Percival J. Parrott, M.A., 
IGlenn W. Herrick, B.S.A., 

Entomologists. 
Hugh Glasgow, Ph.D., 
*Fred Z. Hartzell, M.A. (Fredonia), 

Associate Entomologists. 
RossiTER D. Olmstead, B.S., 
Clarence R. Phipps, B.S., 

Assistant Entomologists. 
Ulysses P. Hedrick, Sc.D., 

Horticulturist. 
fRoLLiNS A. Emerson, Sc.D., 

Geneticist. 
fWiLLiAM H. Chandler, Ph.D., 

Pomologist. 
t Roy D. Anthony., M.S.A., 
*Fred E. Gladwin, B.S. (Fredonia), 
Orrin M. Taylor, 
George H. Howe, B.S.A., 

Associate Horticulturists. 
William C. Stone, M.S., 
Edward H. Francis, M.A., 
Theodore E. Gaty, B.S., 

Assistant Horticulturists. 
Jessie A. Sperry, Director's Secretary. 
Frank E. Newton, 

WiLLARD F. PaTCHIN, 

Lena G. Curtis, 
Mae M. Melvin, 
Maude L. Hogan, 
K. Loraine Horton, 

Clerks and Stenographers. 
Elizabeth Jones, 

Computer and Mailing Clerk. 



Address all correspondence, not to individual members of the staff, but to the 
New York Agricultural Experiment Station, Geneva, N. Y. 

The Bulletins pubUshed by the Station will be sent free to any farmer applying 
for them. 

Connected with Grape Culture Investigations. 
t Members of the faculty of the New York State College of Agriculture affiliated with this 
Station. 

J Resigned to accept position at Pennsylvania State College. 

1 LiBRARV- OF CONGRESS j 



TECHNICAL BULLETIN No. 76. 

ASEXUAL mHEEITANCE IN THE VIOLET 

( Viola odor aid)} 

ROY D. ANTHONY. 

SUMMARY 

In the improvement of fruit varieties the question of fixity of type 
in asexual propagation is of very considerable importance. The use 
of any of the tree fruits in a study of this problem would obviously 
extend the experiment far past the activity of a single investigator. 
In order to hasten work on this question the double violet, Marie 
Louise, which is propagated asexually, was used in a study of the 
effect of selection upon the length of blossom stem. Observations 
were also made of the inheritance of high and low yield. 

Four selection groups were made : long-stem plants of high yield, 
long-stem plants of low yield, short-stem plants of high yield, and 
short-stem plants of low yield. 

The first year it was not realized how important a role plant vigor 
would play and so no record of this factor was made. Since then 
the plants have been graded for vigor twice each season. 

One of the greatest difficulties encountered has been to find 
the best methods of showing the year's work and of making the 
selections for the following year. The method of selecting the 
plants for propagation was changed after two years. 

Two sets of charts were used, one where the yield and stem- 
length of individual plants or of clonal groups are correlated with 
vigor and a second series showing the influence of location in the 
house upon yield, stem-length, and vigor. The length of all blossom 
stems is reported in one-half inch units and this unit is employed 
in the cjiarts. 

Correlation tables are given for the three factors for the entire 
house for each year of the experiment and for each of the four 
selection groups for the last year. 

For purposes of comparison, the four selection groups were reduced 
to the same vigor by means of the regression coefficient. 

Environmental factors caused considerable variation within the 
same greenhouse, especially the first year of the experiment. 

The second year, the vigor and yield were approximately the 
same for the long-high and short-high groups but there was a lower 
stem-length average in the short-stem than in the long-stem selection. 

1 Also presented to the faculty of the Graduate School of Cornell University as a 
thesisin partial fulfillment of the requirements for the degreeof doctcrof philosophy. 

3 



This second year four plants were grown from every plant selected 
as a parent the first year. The record of these four plants showed 
that many of the first selections, based on the performance of one 
plant and without a knowledge of its vigor, were not correct. 

The third year the two high-yielding selections gave a sUghtly 
higher yield than the low selections but the difference was less 
than twice the probable error. At the same time in the two high- 
yielding selections the difference in stem-length in favor of the long- 
stem group was about five times the probable error and in the low- 
yielding selections about nine times. 

The fourth year there were supposed to be sixty-four plants from 
a common parent in 1914. In the long-stem selections, the high- 
yielding group averaged 3.108 ± 0.441 blossoms more than the 
low-yielding plants. In the short-stem selection the high-yielding 
plants averaged 5.787 ± 0.478 more than the low-yielding plants. 
In both the high-yielding and low-yielding selections the long-stem 
plants averaged, respectively, 0.361 ± 0.036 units and 0.495 ± 0.041 
units longer than the short-stem plants. 

The fifth year of the experiment, in the long-stem selections the 
high-yielding plants averaged 1.847 =t 0.754 blossoms more than 
the low-yielding plants. In the short-stem selections the high- 
yielding plants showed a gain of 4.970 =b 0.767 blossoms. The 
long-stem groups in both the high and low yielding selections 
gained 0.368 ± .046 and 0.346 ± .048 units, respectively. 

The process of selection has really been one of isolation whereby 
certain clonal lines have been selected out of the miscellaneous 
population purchased in 1914. We seemingly have proved only the 
existence of asexually inherited differences which probably were 
present before the experiment was begun. No attempt has been 
made to find when or how such differences arose. 

Tho the existence of such differences in the violet makes 
it seem more probable that there may be differences within a single 
variety of any fruit, the labor and the technical difficulties involved 
render it inadvisable for a nurseryman to attempt to find beneficial 
variations among fruits by bud selection. ' 

INTRODUCTION 

THE PROBLEM 

All commercial fruits of America are propagated by asexual means. 
This fact creates a fundamental difference in the problem of the im- 
provement of our present fruit varieties as compared with the improve- 
ment of most vegetables and field crops where reproduction is by 
sexual means. 

The development of the idea of pedigree and all it involves has 
had such a profound influence upon the Hve-stock industry that it 



is only natural for the fruit-grower to be influenced in his thought of 
his trees by this same idea. The growth of this trend of thought 
has been hastened by the exploitation of the term "pedigree" by a 
number of nurserymen who have sought by its use to convince the 
buyer that their trees were better than ordinary trees, tho the 
exact grounds upon which tliis statement was based seem frequently 
to be uncertain even in the minds of the niu-serymen themselves. 

Since 1896, the Geneva Station has been working on a problem 
in orchard fertilization where the question of the fixity of type in 
asexual propagation is of considerable importance. That year an 
orchard of Ben Davis trees was planted for a fertihzer experiment. 
The trees were then top-worked to Rome, the buds all coming from 
a single tree. In 1912 a problem in selection within a clonal fine 
was started when buds were taken 'from the highest and from the 
lowest yielding trees in each fertihzer block and budded on own- 
rooted Spy stocks. These were planted on a uniform soil and are 
now nearly ready to begin fruiting. 

At about the time this second test was started, an attempt was 
made to study the fixity of type in the Baldwin by planting an orchard 
of trees secured from every part of the country and showing as wide 
a diversity as possible in their "pedigree." It is too early yet to 
know what the value of this last experiment will be, but owing to 
the conditions surrounding the orchard it is doubtful if it can be 
relied upon to show conclusive results. 

It is obvious that work of this sort with species as slow in coming 
to matm'ity as are the tree fruits, must, of necessity, run far past 
the span of the working hfe of a single investigator. Even with the 
bush and small fruits progress would still be slow and these fruits 
are very susceptible to environmental changes. Therefore, in order 
to hasten the work on this problem, it was decided, in the spring of 
1914, to grow the double violet, Marie Louise, in the greenhouse 
and to study the effect of selection upon the length of stem of the 
blossoms. That fall eight hundred plants were purchased from a 
commercial grower and planted in the greenhouse. 

The first question to be answered was whether from a mixed popu- 
lation types, or strains, could be isolated which would hold true to 
their selection year after year. The occurrence of "sports" in many 
horticultural crops is weU known, altho when we consider the oppor- 
tunities for their production the number that have been isolated and 
proved to reproduce themselves is almost neghgible. Some recent 
investigations would seem to indicate that citrus is an exception and 
that this genus is in a state of change, producing new types frequently. 

One variety of apple commonly grown in New York, the Twenty 
Ounce, has rather recently produced three sports, while a fourth 
can probably be credited to it. It is doubtful, however, if this 
number of sports has been found in all the other fruits grown in this 
State. Of course such sports represent sufficiently great changes to 



be seen readily and to stand out from the minor fluctuations due to 
environment. Whether or not there are heritable differences too 
small to be detected in the commercial plantations is a question of 
fundamental importance tp all engaged in the attempt to improve 
our fruits. 

As the first five years' work with the violet seems to throw some 
light on this subject and as this year marks the beginning of the 
second phase of the problem, namely, whether by further selection 
isolated types may be shifted in either direction or even spht into 
a multipHcity of types, it has seemed best to publish the data at 
this time. 

LITEKATURE 

So many excellent discussions .of the general subject of bud varia- 
tion have appeared in the last few years that it would seem unneces- 
sary to present a very extensive bibhographic review. However, 
attention should be called to certain of these publications. 

The potato was the first asexually propagated plant in which 
improvement was generally sought by means of selection. A review 
of much of the early work with this plant is presented by Stuart 
(1915).2 

The results of one of the most successful experiments in the selec- 
tion of somatic variations were presented by Stout (1915). His 
work with Coleus did much to call the attention of investigators to 
this problem. 

Dorsey (1916) has given us an excellent review of the literature 
bearing on several phases of the question of bud variation. 

Jennings' work (1916) with Difflugia is very interesting as it in- 
volves a quite different type of reproduction. The chapter devoted 
to bud selection in Babcock and Clausen's book (1918) is a splendid 
critical study of the different phases of this problem. 

The most recent contributions from the horticultural standpoint 
have been Shamel's articles (1918) dealing with citrus fruit improve- 
ment. 

DESCRIPTION OF MATERIAL USED 

The double violet, Marie Louise, for fifty years has been one of 
the most widely grown varieties in commercial and amateur houses. 
It is entirely sterile and is propagated by the separation of shoots 
from the original plant. So far as we have been able to find, the 
variety runs very uniform. While one or two so-called strains have 
been produced, the fact that they have never made their way into 
commercial culture would seem to indicate that they differed little if 
at all from the true variety. Alt ho the previous treatment of the 
800 plants with which the experiment was started was not known, it 
is safe to assume that they would trace back to a very few parent 

^ Reference to Literature Cited, page 30. 



plants in a few generations as nurserymen are constantly propa- 
gating from their best plants and each healthy plant yields eight or 
ten cuttings. 

These plants were placed in the south house of a range of three 
running east from the main house. The south-east corner of the 
house is exposed to some of our coldest winds and fluctuations of 
several degrees in temperature have been noted between this corner 
and the sheltered north-west corner. The main house at the west 
not only shelters from the wind but also has a tendency to decrease 
the Hght received in the north-west quarter of the house. This 
point will be discussed somewhat in detail later. Within the house 
are two tile-bottom benches, each holding 400 plants in fifty rows of 
eight plants to the row. These two benches will be referred to as 
the north and south benches, respectively. 

New plants are propagated each year either by allowing roots to 
form on the shoots, which are put out at the base of the crown, before 
the cuttings are taken, or the shoots are taken off just as the roots 
start to form. The shoots are then put in flats filled with sand. 
When root-growth is well started, the shoots are transferred to thumb 
pots and later, to larger pots and placed in cold frames for the summer. 
The plants are set in the benches about the first of September. 

The first few blooms that form usually have very short stems 
and are imperfect and these are discarded without measuring. The 
first regular picking is made about the middle of October. With a 
slight jerk the stems separate readily from the plant and the entire 
length from the base of the blossom is measured in units of one- 
half inch. 

As it was necessary to tabulate the results before plants could be 
selected for propagation for the following year, the last harvest 
record was usually made early in March, at a time when the plants 
were giving nearly maximum yields. 

METHODS 

House records. — Tho the experiment was to deal primarily with 
blossom-stem length, it was decided to study inheritance of high 
and low yield also as yield records had to be taken in getting stem- 
lengths. As plants were selected for long and short stem and high 
and low yield, this gave four selection groups: long-stem plants 
of high yield, long-stem plants of low yield, short-stem plants of 
high yield, and short-stem plants of low yield. In order to shorten 
the records and the discussion, these groups are usually referred to as 
follows : long-high, long-low, short-high, and short-low, respectively. 

The records are taken in the greenhouse on large sheets 14 by 17 
inches, cross-ruled into quarter-inch squares. The plant number is 
placed at the left . and the date at the top. Each flower-stem is 
measured and the length to the nearest one-half inch placed in the 



next square. When each harvest is over, a vertical line is drawn 
outside of the last record of the highest yielding plant. In this way 
the first records of the following harvest for all plants are in the same 
vertical Une of squares, making it easy to study the yield fluctuation 
of any particular plant from harvest to harvest, and showing the 
relation of any plant to the maximum yield of that harvest. 

With conditions at their best, the bed is picked over every week or 
ten days but in very cold weather it may be three weeks or more 
between harvests. When the large record sheets are filled they are 
totaled for each plant. In an average season, when the third series 
of sheets is filled, the records are discontinued and propagation com- 
menced for the following year. For the first year the various factors 
were studied for each of these three harvest periods but since then 
only for the total harvest. 

When the experiment was begun it was not reahzed how important 
a role plant vigor would play and so the first year no record was 
made of this factor but the need of it was clearly seen when it became 
necessary to select plants for propagation for the following year. 
There seemed to be no feasible way of weighing the plants or measur- 
ing their leaf surface and so the vigor was estimated in percentage, 
the most vigorous plants being graded as 100 and a few plants, with 
only a half dozen or so leaves and practically no blossoms, graded 
as 10. Two persons cooperated in this work, each checking the 
other, and it is doubtful if a third person would have shifted any of 
the plants more than ten points in the vigor score. The vigor was 
taken twice, once when the plants were coming into full bloom early 
in the winter, and a second time just before the selection of the plants 
for propagation for the next year. In computing final records the 
average of the two observations was used. 

In a very general way, plants recorded as having a certain vigor 
were somewhat alike from year to year; but, because of the varia- 
tions of the plants in diff"erent seasons, it is not possible to compare 
the vigor records of two different years with each other except in so 
far as each is considered as a statement of the relative values of the 
plants for that particular year. 

Statistical methods. — In order to simphfy the problem of studying 
the plants and especially of making selections for the next year's 
propagation, the number, yield, and stem-length of each plant was 
placed on a single card and the cards grouped according to the line 
of selection. These cards could then be shuffled for any factor, and 
frequency charts and correlation tables very quickly prepared. This 
paethod has resulted in a considerable saving in time and has been 
a very important help in analyzing the data. 

One of the greatest difficulties encountered in the experiment was 
to find the best method of showing the results of the year's work 
and of making the selections for the following year. The method 
of selecting the plants for propagation was not the same for the first 



two years as is now used. In these early years the records were 
thrown into frequency tables and certain maximum or minimum 
yields and lengths selected until the number of plants which it was 
planned to propagate was isolated. This is well illustrated by Chart I 
which shows the selection standards for the first year in connection 
with the frequency curves of length and yield. The lines from which 
the arrows are drawn show the selection limits. 



Chart I. — Frequency Curves of Length and Yield, 
1914-15. 



Length. 



Yield. 




Length in one-half inches. 



Yield per plant. 



By selecting the plants from the frequency curve and without knowl- 
edge of their vigor, any plants which were high-yielding because of ab- 
normally high vigor were selected and the same was true with plants 
of low yield, due to lack of vigor. This objection has now been 
overcome by a somewhat different method of selection. Each plant 
or group of plants is plotted according to vigor on a chart with a 
common base line and with the yield plotted above and the stem- 
length below the vigor hne. (See Chart IV.) In this way two 



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12 

points are located for a single plant or group of plants, the one directly- 
above the other, and the line connecting them passes thru the 
vigor value of the plant. As an aid in selection, a straight line was 
drawn thru each population in such a way as to divide the members in 
the various vigor classes into nearly equal parts. Inasmuch as the 
individuals near this hne, the average plants, were discarded when- 
ever possible, tliis method of dividing the population was deemed 
sufficiently accurate. To check this point, however, in several cases 
the straight line was determined by the formula, y=mx + n. The 
two fines were found to differ but little. Values lying above the fine 
would then represent high-yielding or long-stem plants and those 
below the fine low-yielding or short-stem plants. 

By selecting from this chart it was possible to find those plants 
which were above or below the average at any particular vigor. 
This method has been used the last two years with very satis- 
factory results. In 1915-16 the selections were made on the basis 
of the average performance of the four plants tracing from a common 
origin the previous year. The following year the average of the six- 
teen plaiTts with common origin v/as used and so on for each year, the 
total number tracing from a single plant in 1914 increasing by mul- 
tiples of four each year. 

A second series of charts (Chart III) is used to show the influence 
of location in the house upon the three factors studied and the rela- 
tive fluctuation of those factors. These charts are also useful in 
showing the sudden jumps that frequently occur in passing from one 
type of selection to another when the 'plants lie in adjacent rows. 

Correlation tables are made for the three factors for the entire 
house and for each of the four selection groups, and for each of these 
the following values are computed : Coefficient of correlation of 
vigor and length, vigor and yield, and length and yield ; the standard 
deviation of length, yield, and vigor; the mean values of length, 
yield, and vigor; regression coefficients; and the corrected means 
where yield and length have been reduced to a common vigor by 
the regression coefficients. 

These values are summarized in Table I and in the appendix 
are placed the correlation tables for the entire house for each year 
of the experiment and the correlations for each selection group for 
the last year. To include all the correlation tables seemed unneces- 
sary since those which were selected show the general trend of the 
correlations. 

A study of the correlation tables themselves without reference to 
computed values is of considerable help as it shows roughly the 
degree of correlation, the approximate averages, and the presence 
of any abnormal plants which need further study. If any sporting 
occurs in the factors studied it can be quickly detected by this means. 

In studying the 1915-16 records a number of the correlations 
were worked out for the first and for the third periods of harvest. 



13 



The correlation between vigor and yield in the first group of pick- 
ings was low but for the third group considerably higher except for 
the short-low group and in the total this correlation is still higher 

with all groups. A study of the formula for correlation, r2/„=- — —", 

shows the mathematical reason for this. The total yields for the 
third group were of course considerably higher than for the first 
group of pickings and the total for the three groups very much higher. 
Thus, we have a constantly increasing ^dd in the numerator, 
while in the denominator the standard deviation for vigor does not 
fluctuate materially and the standard deviation for yield does not 
increase as rapidly as the summation so that we have a constantly 
increasing value for the coefficient of correlation. The fundamental 
reason is probably that by the third period of harvest the plants have 
reached their maximum production and under such conditions prob- 
ably show greater correlation than earlier in the season. This varia- 
tion in the correlation is another reason why it is difficult to compare 
different years with each other. 

In Table II are summarized for the last four years the corrected 
mean yield and mean stem-length for the house and the differences 
for the contrasted selection groups. Such a summ.ary is very con- 
venient in the analysis of the data. 



Table II. — Four Year Summaries of Yields and Stem-lengths. 

Yield and length for each selection group computed for each year to average vigor 

of house. 



Yields. 


Stem-lengths. 


Group yields. 


Long stem. 


Short stem. 


Group 
lengths. 


High yield. 


Low yield. 


1915-16 

High yield 

Low yield 


34.08 ±.488 
33.65 ±.631 


32.18± .659 
34.36± .691 


Long stem. . 
Short stem. . 

Long stem. . 
Short stem. . 

Long stem. . 
Short stem.. 

Long stem . . 
Short stem. . 


8.920±.029 
8.537±.G40 


8.483±.043 
8.537±.043 


Gain or loss. 

1916-17 

High yield 

Low yield 


.43 ±.797 

31. 893 ±.537 
30. 460 ±.605 


— 2.18± .954 

32. 837 ±.603 
31. 436 ±.672 


.383 ±.049 

«.635±.028 
8.437±.034 


— .054±.060 

8.603±.033 

8.243±.028 


Gain 

1917-18 

High yield 

Low yield 


1.433±.808 

17. 695 ±.312 
14. 587 ±.311 


1.401 ±.902 

19. 903 ±.393 
14.116±.271 


.198 ±.044 

7.758±.026 
7.397±.025 


.360 ±.044 

7.812±.033 
7.317±.024 


Gain 

1918-19 

High yield .... 
Low yield 


3.108±.441 

36. 484 ±.505 
34. 637 ±.572 


5.787±.478 

40. 359 ±.569 
35. 389 ±.515 


.361 ±.036 

8.409±.030 
8.041±.036 


.495 ±.041 

8.525±.031 
8.179±.032 


Gain 


1.847 ±.754 


4.970±.767 


.368 ±.046 


.346 ±.048 



14 

More should be said of the use of the regression coefficient. A 
study of the coefficients of correlation shows the high degree of 
correlation between vigor and yield and vigor and stem-length. For 
this reason it became necessary to find some way of reducing the 
various groups to a common vigor in order to compare yield and 
stem-length. Tliis can be done with sufficient accuracy by means 
of the regression coefficient. ^ The four selection groups were reduced 
to the same vigor as the house average by multiplying the difference 
between the group average vigor and the house average vigor by 
the regression coefficient. If the group vigor was less than the house 
average this value was added to the group average length; if the 
group vigor was greater than the house average, it was subtracted 
from the group length. 

INFLUENCE OF LOCATION IN HOUSE UPON YIELD AND 

STEM-LENGTH 

On first thought, one would expect that the house conditions were 
sufficiently uniform so that environmental variations could be 
excluded but a study of the year's record shows this not to be the case. 
Even after five years' study all of the factors causing variation can 
not be stated positively but the two most influential are probably 
temperature and fight. As both of these differ in different seasons 
we find, in studying the house from year to year, indications of yearly 
variations so that any conclusions drawn from the first year's record 
must be applied to any other year only with many reservations. 

Chart II shows the yield and average stem-length for 1914-15 
for groups of sixteen plants in two adjacent rows as they stood in 
the two benches. Care was taken to have the soil and cultural 
treatment as uniform as possible and so it seems safe to say that 
any fluctuations of the groups are due to the influence of location 
in the house upon plant growth. In other years variations may be 
due to the selection as well as to the influence of position. 

On the chart we notice a fluctuation in the north bench which 
has appeared to a greater or less extent in every year and has been 
an important factor in compficating the results in this bench. In 
the west half of the bench (the left side of the chart), the yield shows 
a peak extending about one-third of the way down the bench and 
there is a remarkable rise in the stem-length of this area. Year 
after year in this part of the bencii have been the most vigorous 
plants, those with the longest stems, and, usually, those with the 
highest yield in the entire house. The eastern haK of the bench 
this first year shows a constantly increasing yield and stem-length. 
This increase does not seem to be as pronounced ,in other years and 
just why it should show to the extent that it does this first year is 

^ The formula used was: Regression coefficient of length to vigor = Tlv • 



15 



Chart II. — Influence of Location in House upon Yield and Stem-length, 1914-15. 

Length Yield 

in 2 per 
inches plant. 



9.10 35 



8.90 30 



.70 25 



.50 20 



.30 15 

Rows 




North bench. 



V Stem-length 



Yield 



10 



30 



9.10 35 



.90 30 



8.70 25 



.50 20 



.30 




South bench. 



15 

Rows 



Yield 



\ Stem-length 



10. 



20 



30 



40 



50 



16 



not known. In the south bench there are some rather sharp fluctua- 
tions in stem-length and, in general, the yield fluctuates with the 
length. Taking the whole of the south bench into consideration 
no marked change seems to show which could be laid to environment. 
In the Hght of the other years' records, however, the tendency of 
the stem-length to increase slightly at the west end while the yield 
decreases slightly is significant, and it is probably safe to say that 
the western half of the bench is somewhat more favorably located 
for stem-length development than the eastern half. 

GENERAL DISCUSSION OF RESULTS FOR 1914-15 

The first year no record was taken of vigor, only the total yield 
of each plant and the stem-length of each blossom being recorded. 
Altho the lack of a record of vigor decreases the value of the first 
year's results to some extent, our knowledge of the general relations 
of stem-length and yield gathered from the other four years enables 
us to interpret the results for the first year with considerable accuracy. 

House-values. — Because of the lack of the vigor record, it was pos- 
sible to correlate only length and yield. The coefficient of correla- 
tion for the entire house was 0.415 ± .019, very considerably lower 
than the following year when it was 0.729 ± .012. This low corre- 
lation is due to the failure of the two factors to respond in the same 
degree to the house conditions affecting the west end of the two 
benches. This condition seems more noticeable this year than in 
the following years. The average length for the house was high, 
8.836 ± .012 units, in proportion to the yield of 27.969 ± .259 
blooms, as shown in Table III. 

Table III. — Fn^E Year Summaries. 
Mean values for entire house. 



Year. 


Yield. 


Length. 


Vigor. 


1914-15 


27. 969 ±.259 
35.590±.306 
31.747±.306 
16.368±.176 
36. 367 ±.278 


8.836±.012 
8.600±.018 
8.489±.016 
7.535±.015 
8.3S0±.016 




1915-16 


49. 996 ± 505 


1916-17 


53 177± 471 


1917-18 


53. 591 ±.567 


1918-19 


81.447±.339 







It is probable that the commercial grower from whom these plants 
were purchased had been selecting for high vigor for years, and thus 
had been unconsciously selecting for long stems and high yield. * 

Selection of plants for 1915-16. — The method used in selecting the 
plants for propagation during the first years has already been de- 
scribed. Because of the errors involved in this method and because 



17 

of lack of knowledge of the vigor of the plants, many were marked 
for propagation along the four hnes of selection which, in the light 
of our present knowledge, should have been discarded or should 
have been included in a different class from the one in which they 
were placed. 

GENERAL DISCUSSION OF RESULTS FOR 1915-16 

Before it was realized to what extent the factors under observa- 
tion would be influenced by house conditions, the plants for the 
1915-16 crop were planted in the house. The long-high and short- 
high groups were placed in the south bench and the long-low and 
short-low groups in the north bench. As the total number- of plants 
this year was kept to about 700, all the groups save the long-high 
occupied somewhat less than a quarter of a bench. 

House values. — From Chart III, where each point represents the 
average for twelve adjacent plants, it will be seen that, on the south 
bench, the vigor and yield were approximately the same for the 
two groups but that the line of stem-length is somewhat lower in 
the short-stem selection than in the long-stem selection. 

On the north bench the region of high yield discussed under the 
previous year's data does not begin until the seventh or eighth 
row but seems to carry to the east to about the thirty-second row, 
covering practically the entire bench so far as it is occupied by these 
two groups. In this bench the fluctuations in vigor make it impos- 
sible to observe any certain differences in stem-length or yield in 
the two selections. Referring to Table II, where the summaries for 
the four years are given, we note that the average stem-lengths for 
these two groups do not show a significant difference whfle the stem- 
length difference in the south bench is .383 ± .049 units in favor 
of the long-stem selection. Owing to the differences between the 
two benches it is difficult to draw any conclusion with regard to 
yield. On the west ends of the two benches the high-yield is sHghtly 
better than the low-yield selection when reduced to a common vigor. 
Most of the short-low plants on the north bench lie in a more favor- 
able position than the short-high plants on the south bench and this 
probably accounts for this group being the higher yielding one 
contrary to the selection. 

Selection of plants for 1916-17. — Each half row in the .year 1915- 
16 originated from a single plant of the previous year. At the time 
that that single plant was selected to act as a parent only the one 
year's record was available to use in judging in which group the plant 
should faff. At the end of 1915-16 the record of the four plants 
with a common parentage showed clearly that a number of groups 
did not belong in the classes in which they had been placed. Some 
indeed were so far out of the class in which they were first placed 
that it seemed wise to select them for continued propagation in one 



18 



Chakt III — Influence of Location in the House on Yield, Stem-length, and Vigor, 

1915-16. 




North bench. 
* Yield. 



Stem-length. 
Vigor. 



30 40 

Short stem low yield. 



South bench 



Long stem high yield. 



30 40 50 

Short stem high yield. 



20 



Ave. length 
in i inches. 



Chart TV. — Relation ( 
Record o 



■ : i < 



30 



40 






» 



Chart IV. — Relation of Yield and Stem-length to Vigor, 1915-16, 
Record of four plants from same parent. 



Total yield, 
foux plaBts. 



c? Retained for high yield. 
P Retained for low yield. 























t 


230 


















« 




210 
















s 






190 




















i 
« 


170 






t 




*- » 


s' »»»SJ 






'I 


» >i 


150 








': 


,r.r 


«' 






/ - . , 








V 


A 


J*' 




*• 


J" 


t 




130 






- 


r^' 


s''- 




i 




» 




110 




* *• 




« < 


















» « 


*^ 


« 


H 














■< % 


" 


K- 


t 


i 










90 




« 




« 














70 
igor 




rf 


















20 


30 




40 




50 




60 




70 80 90 


9.75 


■ 










S 






8 


t 


9.50 










S 




«* 


« 


J 


t 


9.25 










ir 


i 








« 








1 




s 


«s. .. 


K 




v*». ?» 


* s 


9.00 






K 


a< 




•1* 


}. 




a 


« 


8.75 














II 






. « 








"" 


W 


nH 




K 




s 








» ■« 


y 


% 


* » 










8.50 




» 


% 


•$' 


* 






»l 












r' '^ 


•«? 


9 J? 










8.25 




* 






t 


< » 

' 1 


Ir 








8.00 


« 


V 


«-*■ 


MJ 












c? Retained for long stem. 


7.7S 


• 


















P Retained for short stem. 



Ave. length 
in i inches. 



19 

of the other groups in which they now appeared to fall. Others of 
these abnormal plants were discarded. As all these plants were 
included in the group tabulations they have served to smooth out 
the results to some extent and hide any effects due to the selection. 
Chart IV shows the distribution of the groups of four plants. The 
groups selected for further propagation are marked with the arrow, the 
direction of the arrow indicating the direction of the selection. The 
method illustrated in this chart was not developed until later and an 
inspection of the chart shows that some groups were retained which 
should have been discarded while many were discarded which should 
have been retained. 

GENERAL DISCUSSION OF RESULTS FOR 1916-17 

This year there were sixteen plants tracing back to a single plant 
placed in the house in 1914. These sixteen plants were set out in 
two rows of eight each, extending across the bench. Groups repre- 
senting the same type of selection were scattered thru the house 
in such a way that the influence of house variations was probably 
very largely done away with so far as the summaries are concerned. 

From an inspection of the graphs of the values for length, yield, 
and vigor, plotted for position in the house as shown in Chart V, 
it would seem that there was somewhat less fluctuation on the north 
bench than was the case the first two years, especially in regard to 
yield and stem-length. There was, however, a tendency for the 
first sixteen rows from the west to be somewhat better than the 
next sixteen. In the previous year the high area on the north bench 
seemed to extend from about Row 10 thru Row 31. There was 
considerably more vaiation in the south bench than in the previous 
two years, there being an especially high area between Rows 8 and 
18. The different selection classes were so scattered thru the house 
that no one class seems to have been affected by these areas more 
than the others. 

H.ouse values. — Toward the latter part of the season, nematodes 
were found on the roots of several plants. These decreased the 
vigor and the yield of blossoms on a number of plants but in spite 
of this the house average yield was 31.747 ± .306 from November 3, 
1916, to March 5, 1917. (See Table III.) The average stem-length 
for the same period was 8.489 db .016 units, or 4.244 inches. It is 
probable that, because of the nematodes, our vigor standard was 
slightly lower than in the previous year as the average length of 
practically 8.5 units was.from plants with an average vigor of 53.177 
± .471, while in 1915-16 the average length of 8.600 ± .018 was 
from plants with an average vigor of practically 30. 

Group yields. — In the two groups selected for long stems, the 
high-yielding selection gave 1.433 zh .808 blooms more than the low 
selection and in the two short-stem groups the difference between 'the 



20 



Chart V. — Influence of Location in House on Yield, Stem-length, and Vigor, 1916-17. 

O M) 2 

.sp g .a 
> >^ >* . 

70 9.25 50 



65 9.00 45 
60 8.75 40 
55 8.50 35 
50 8.25 30 
45 8.00 25 
40 7.75 20 



I 













\ 




\ 






\ 




.\ 






\ 




,\ 


1 
1 
1 


\ 


/ 




\ 


/X 


\ 




/' 





North bench. 
"~~ • -~~ • Vigor. 

...... Length. 

Yield. 




South bench. 



Chart VI. — Rela 
R( 



Ave. yield 
per plant. 



42 



36 



30 



24' 



^18 
Vigor 



9.00 



8.50 



8.00 



35 



40 



45 



7.50. 
Ave. length. 



Ave. yield 
per plant. 



Chart VI. — Relation ou Yield and Stem-length to Vigor, 1916-17. 
Record of 16 plants from same parent. 



48' 
























42 


















* 
* 


« 




36 










» 










.' r 




30 






• 




W ■ 




< 












24- 




n 


X 


m «• 




t 








r 






















c? Selected high.i 
P Selected low. 




' 18 






Jt 


















Vigor 


33 




40 


45 




50 




55 


GO 


63 

« 


70 


9.00 










1C 


s 




« 








8.50 




M 


« ■■ 


% 

•t X 

n 


K 

< 


•• 


H 


s 
i 








8.00 
7.50 








V 


i 










S Selected long. 
P Selected short. 





Avo. lengtli. 



21 

high and low selection was 1.401 ± .902 blooms. (See Table II.) 
This is not a large difference and in each case it is less than twice the 
probable error of the difference.^ 

Group lengths. — In the two high-yielding selections the average 
stem-length of the short-stem group was .198 ± .044 units below the 
average for the long-stem group and in the two low-yielding selec- 
tions the average stem-length of the short-stem group was .360 ± 
.044 units below the long-stem group. (See Table II.) In the high- 
yielding group this difference is about five times the probable error 
and in the low-yielding group about nine times the probable error. 
It must be borne in mind that the selection, which is really a process 
of isolation, is not acting rapidly enough at the end of the first two 
selection periods for us to expect any very great difference. Such 
differences as these were, therefore, a surprise to all engaged in the 
work and may be considered as very significant in throwing light 
upon the trend of the experiment. 

Selection of plants for 1917-18. — In making the selection for the 
following year, groups tracing back to a common parent plant in 
1914 were treated as units. Chart VI shows the distribution of 
the average values for these groups and those selected for further 
propagation. 

GENERAL DISCUSSION OF RESULTS FOR 1917-18 

It was planned for this year that there should be sixteen rows of 
four plants each, all tracing back to a single plant in the 1914-15 
population. It was not possible in all cases, however, to continue 
propagating at the rate of four to one and so some of the groups fall 
below a total of sixty-four. Plants having a common origin were 
not separated, but those groups which had a common selection were 
scattered through the house in order to eliminate, so far as possible, 
the influence of house variations. It would have been somewhat 
better had the units of sixty-four plants been split up into two or 
four groups and placed in different parts of the house. 

House values. — The very unusual cold, together with the age and 
poor condition of the house, resulted in a very light yield. The 
average yield for the house for a period extending from November 
20, 1917, to March 12, 1918, was 16.368 ± .176 blossoms. The 
average stem-length was 7.535 ± .015. But one vigor record was 
taken this year, that at the end of the picking season. The average 
vigor was 53.591 dz .567. (See Table III.) 

Group yields. — In the long-stem selections the high-yielding group 
averaged 3.108 dz .441 blossoms more than the low-yielding plants.. 
In the short-stem selection the high-yielding plants averaged 5.787 
± .478 more than the low-yielding plants. (See Table II.) 

* The probable error of the difference is found by extracting the square root of the 
sum of the squares of the probable errors of the means which are compared. 



22 

Group lengths. — In the high-yielding selection the long-stem plants 
averaged .361 ± .036 units longer than the short-stem plants, and 
in the low-yielding selection the long-stem plants were, on the aver- 
age, .495 ± .041 units longer. In comparison with the probable 
errors these differences are sufficiently large to remove any doubt 
of their value. (See Table II.) 

Seledi7ig plants for 1918-19. — The general correlations of the three 
factors are shown graphically in Chart VII. Here, also, are indi- 
cated the plants retained for the next year. 

This chart shows even more clearly than the figures given above 
that our process of isolation has separated out distinct groups. Thus, 
in the north-west quarter of the house, the long-low group and the 
short-high group have the positions of the stem-length and yield 
hnes reversed at nearly the same vigor. The yield line of the long- 
high group is much higher at lower vigors than in the short-low group. 
On the west end of the south bench, the two contrasted selections, 
long-low and short-liigh, show a complete reversal in the relative 
positions of the yield and length lines. 

In Chart VIII the average yield and average stem-length of 
groups of twelve adjacent plants are plotted to the average vigor. 
Since there were as high as sixty plants in a single hne, the points 
on the chart do not now represent the entire performance of a partic- 
ular selection as has previously been the case in this type of chart. 

GENERAL DISCUSSION OF RESULTS FOR 1918-19 

Propagating the selected plants at the rate of four to one, there 
should be 256 tracing back to a common origin in 1914. 0\^ing to 
the vicissitudes of propagation, however, this number was not 
retained in every case. In one or two cases the number was very 
considerably cut down, but in such cases this was done largely 
because of lack of room or the uncertainty as to whether that par- 
ticular Hne would be continued further. The larger populations 
were broken into three groups and placed in different parts of the 
house. In this way it is probable that whatever house fluctuations 
there were have not seriously influenced the final average results. 
In general, the house fluctuations this year were not extreme. In 
Chart IX, the usual peak is seen on the north bench but it is not 
extreme and seems to affect yield more than the other factors. The 
south bench shows many minor fluctuations but, on the average, 
it is fairly uniform. 

House values. — The total yield per plant from October 18, 1918, 
to January 31, 1919, was 36.367 ± .278. The average stem-length 
was 8.33 ± .016. (See Table III.) The first two harvests of the 
season were very heavy and, as is always the case with the first 
harvest, there was a high proportion of short-stem blossoms. In 
other years the first harvest has been discarded because of these 



Chart VII. — Influence of Location in House on Yield, Stem-length, and Vigor, 1917-18. 

North bench. 



b bo 3 



> Hq tX 




90 8.75 27 J 


\ 


SO 50 24 


\ 




'.\ 


70 8.25 21 


\\ 


60 8.00 18 


\ 1 


50 7.75 15 


\ 






40 7.50 12 




30 7.25 9 

20 7.00 6 
Rows 








CD 



5,//. 



<® 



/ 



&> 



NOKTH SIDE 

Yield. 

Stem-length. 

Vigor. 

CD Re 

tained for 
1918-19. 



10 



20 



30 



40 



50 



80 8.50 24 



CO 8.00 18 



40 7.50 12 



20 7.00 6 
Rows 




&i. 



10 



L.^. ^ 




South side 



30 



40 



50 



24 



o 

W) 

> 




a 
►3 


3 


90 


8 


75 


27 


SO 


S 


50 


24 


70 


8 


25 


21 


60 


8 


00 


18 


50 


7 


75 


15 


40 


7 


50 


12 


30 


7 


25 


9 




7 


00 


6 






Rows 



Chart VII {continued). 
South bench. 




t-.L, 



\ 



\ 



LM 



^Ji 



North side. 
Yield. 

Stem-length. 
Vigor. 



10 



40 



8.50 24 



60 8.00 IS 



40 7.50 12 



20 7.00 6 
Rows 




'(& 




\ .' S.4. 



xy 



South side. 



50 



Ave. yield 
per plant. 



25 



15 



10 



Chart VIII. — Rela 



"t 



-19. 



ined 
^0. 



5^^ . 

Vigor I 20 



30 



40 



.50 



8.00 



7.50 



1 * 



I « 



7.001 
Ave. length 



Ave. yield 
per plant. 



25 



20 



Chart VIII.— Relation of Yield and Stem-length to Vigor, 1917-18. 
Record of 12 adjacent plants. 

t 



t 



"\ « 



"9 



<S Selected for high yield. 
P Selected for low yield. 



Vigor 



* t 



% a' 



8.00 



7.50 






* « • • I 



7.00 
Ave. length 



.« . 



<S Selected for long stem, 
p Selected for short stem. 



2^1 



CHA.ET IX. — Influence of Location in House on Yield, Stem-length, and Vigor, 191S-19. 
t- Averages of 8 plants. 



95 9.00 45 



90 8.50 42 



85 8.00 39 



80 7.50 36 



75 7.00 33 



70 6.50 30 



North bench. 



65 


6.00 27 




Rows 


95 


9.00 45 


90 


8.50 42 


85 


8.00 39 


80 


7.50 36 


75 


7.00 33 


70 


6.50 30 


65 


6.00 27 




Rows 




North side. 

Yield. 

Stem-length. 

Vigor. 

<2iX^= Retained 
for 1919-10. . 



<S5 



(i3 0> 



10 



74 F 



20 
90 B 



30 



40 



51 D 




^.i- 



(© 



.// 



10 



74 F 



20 
50 E 



30 
39 H 



40 



26 





^ 


D.+J 












-ii ^ 


W) 




.ii'p. 


> 


h-I 


|x 


95 


9.00 45 



Chart IX {Continued). 
South bench. 



90 8.50 42 



85 S.OO 39 



SO 7.50 36 



75 7.00 33 



70 6.50 30 



65 6.00 27 



60 5.50 24 
Rows 



95 9.00 45 



90 8.50 42 



85 8.00 39 



80 7.50 36 



North side. 
Yield. 

— . — — — - Stem-length. 

<4ll) = Re- 
tained for 
1919-20. 




50 E 



75 


7.00 33 


70 


6.50 30 


65 


6.00 27 


60 


5.50 24 
Rows 




10 



90 B 



20 
61 D 



30 



40 



50 E 



Chart X.— Relation 
Ri 



Ave. yield- 
per plant. 



63 



8 * 
i 

it 



70 



75 



2S i 



I 2 



ms 
■ds. 
ob- 

lOf 

ery 
lits 

In 
ich 
ear 
;his 

50 
ing 

,nts 
ats. 
i,me 
red 
; to 
•nly 
3ne 
lec- 
rom 
ove 
ned 
ort- 
3ms 
d is 
ibly 

mts 
ort- 
.346 
reat 
the 
s in 

lines 
larts 
3e in 
the 
■the 
raph 
'rom 
n or 
ases, 
sible 



6.80 

Ave. length 
in 5 inches. 



Chaet X.— Relation of Yield and Stem-length to Vigor, 1918-19. 
Records of 8 adjacent plants. 



Ave. yield 
per plant. 



48 



45 



42 



39 



Ave. length 
in i inches. 



* * e ** 



^ h J s * « . 

t i I ^ < 

* » * t 



aa 






i 




i 


30 












27 




t 




St 




24 












Vigor 


63 






70 




8.90 












8.60 








a 




8.30 










» 


8.00 












7.70 






8 


i 




7.40 




i 




• 




7.10 












6.80 








s 








1 f 



j?e 



n 



8 » 



X 



i 



J 



« J i i r I 









t * t „« ? 

t i 



t .i 



Yield. 



c^High. 
iPLow. 



95 



Length. 



c?Long. 
P Short. 



27 

short-stem blossoms but, on account of the large number of blooms 
in the first harvest this year, it was decided to retain it in the records. 
Because of including these short stems, the average length has prob- 
ably been decreased several tenths of a unit. In the third group of 
harvests, that extending from December 24 to January 31, a very 
considerable proportion of the plants averaged well above ten units 
in length. The average vigor this year was 81.447 ± .339. In 
other years it has been the custom to arrange the vigor scale in such 
a way that the average would be approximately 50, but this year 
such a large proportion of the plants showed extreme vigor that this 
did not seem advisable. Plants recorded as having a vigor of 50 
per cent this year were probably fairly comparable with those having 
a similar vigor record last year. 

Group yields. — In the long-stem selections, the high-yielding plants 
averaged 1.847 ± .754 blooms more than the low-yielding plants. 
(See Table II.) This is less than the difference between the same 
groups in the previous year. This fact may probably be considered 
as another indication that our process of selection is beginning to 
isolate distinct groups because this year there are progeny of only 
two of the original plants represented in the long-high group. One 
of these plants, 51 D, which is the clon retained to continue the selec- 
tion for another year, is not strictly a high-yielding clon, but, from 
its performance of other j^ears, would be graded as somewhat above 
the average. It is retained, however, because it is long-stemmed 
and is sufficiently high-yielding to answer the purpose. In the short- 
stem selection the high-yielding plants gave 4.970 ± .767 blossoms 
more than the low-yielding plants. Tliis great difference in yield is 
due to the fact that 90 B, the short-high strain, is a remarkably 
high-yielding group. 

Group lengths. — In the high-yielding strains the long-stem plants 
were .368 ± .046 units longer in blossom stem-length than the short- 
stem plants and, in the low-yielding strain, the difference was .346 
rb .048. (See Table II.) These differences are not quite as great 
as in the previous year but, as before explained, this is due to the 
inclusion in the records of a large number of short-stem blooms in 
the first picking which in previous years were discarded. 

When the house was planted in the fall of 1918 the different lines 
were broken into groups, the groups being placed in different parts 
of the house. In all there were twelve groups. A study of these in 
Chart IX shows that in seven cases the relative position of the 
graph of both stem-length and yield indicates clearly the type of the 
selection of that group. In each of the other five cases, the graph 
of either stem-length or yield is in the position to be expected from 
its types of selection while the other graph is either uncertain or 
apparently opposite to the selection. In two of these latter cases, 
the two groups of 74 F on the west end of the north bench, a possible 



28 

cause for the failiire to follow the selection may be suggested. If 
we refer back to Chart II we see that with the mixed population of 
1914-15, in the area occupied by these two groups, there was a dif- 
ferent relation between the stem-length and yield lines than in the 
rest of the house. As has been suggested before, this difference is 
probably due to a shght change in the environmental factors in the 
northwest corner of the house. 

The way in which the plants adhere to their respective types of 
selection is also shown in Chart X. Here the whole population is 
shown in groups of eight plants and the character of the selection of 
each group is graphically indicated. It is remarkable the small 
number of groups which fall in areas occupied by groups of the 
opposite selection. 

CONCLUSION 

The first five years of selection have isolated four well-defined 
groups. The differences between the opposite selections are suffi- 
ciently large to show graphically on the charts of the two benches 
for the fourth and fifth years and, in comparison with their probable 
errors, make it certain that they are not due to chance variations. 
It is believed that variations due to lack of uniformity of conditions 
within the house have been so nearly eliminated that the final results 
are not materially affected. 

When this work was started it was the opinion of those connected 
with it that such a result as this would not be secured and probably 
most pomologists would have held the same opinion. For this 
reason there is justification in restating certain points and showing 
the possible apphcation of the results. 

During the five years reported in this publication, the process of 
selection has really been one of isolation whereby certain clonal hnes 
have been selected out of the miscellaneous population purchased in 
1914. In nearly every case each plant within the clon has been the 
parent of four plants used the following year. It follows, then, 
that we seemingly have proved only the existence of asexualiy inheri- 
ted differences which probably were present before the experiment 
was begun. No attempt has been made to find when or how such 
differences arise. 

That differences have been found in the violet which could be 
passed on from bud generation to bud generation does not prove 
that similar differences may be found in the apple but it does make 
it seem more probable that such differences exist. Unfortunately, 
from the standpoint of practical application, the labor and the tech- 
nical difficulties involved in proving that an observed difference is 
really transmissable and not simply a temporary response to an 
environmental change make it seem inadvisable for a nurseryman to 
attempt such a problem. 



29 

The four selection groups now contain only five pure lines, each 
tracing back to a single plant in 1914. Whether these clonal hnes 
are pure hnes in the sense that Johannsen has applied that term 
or whether they are simply stages beyond which we will go to greater 
differences, thru further selection, is now the problem. 

PLANS FOR CONTINUING THE EXPERIMENT 

There had been no selection within any clonal line up to the spring 
of 1919. At that time it was decided that fifty parent plants would 
be saved in each of the four types of selection as this would give the 
800 plants necessary to fill the house. In all but one type these 
fifty were selected from one clon only. As there were from 100 to 
150 plants from which to select the fifty parent plants this afforded 
an opportunity to choose those plants which best answered the 
conditions of their particular type. Instead of selecting single 
plants, groups of four, each tracing back to a single parent the pre- 
vious year, were selected. 

In one group, the short-high selection, two plants were used to 
start two new lines in an attempt to break up a clon by selection 
within it. One of these plants was selected for long stem and the 
other for short stem. In choosing these, care was taken to find two 
plants which did not have a common origin until the original 1914 
plant was reached. These two lines will be continued with intensive 
selection. 

These general plans will be followed for another five years to test 
the fixity of the clonal lines. 

ACKNOWLEDGEMENTS 

Thruout the five years of the experiment Mr. Joseph Welling- 
ton has had the oversight of the propagation and the harvesting. 
This has involved a great deal of work and no small part of whatever 
value the work has is due to the care and accuracy with which the 
records were taken in the greenhouse. He has also assisted in the 
preparation of the data for pubhcation. 

The author is greatly indebted to Dr. H. H. Love of the Depart- 
ment of Plant Breeding of the New York State College of Agriculture 
for many helpful suggestions and advice in the biometrical work. 

The initiation of this work was due to Dr. U. P. Hedrick. For 
many years he had been interested in the question of pedigree in 
fruit and saw the possibilities in extending this study to some plant 
where results could be secured more rapidly than with the apple. 
His direction has been one of the factors in the success of the work. 



30 



LITERATURE CITED 

Stuart, Wm. 1915. Potato breeding and selection. U. S. D. A. 
Bui. 195. 

Stout, A. B. 1915. The estabKshment of varieties in Coleus by the 
selection of somatic variations. Carnegie Inst. Wash. Pub. 218. 

Dorsey, M. J. 1916. The inheritance and permanence of clonal 
varieties. Proc. Am. Soc. Hort. Sci. 1916. 

Jennings, H. S. 1916. Heredity, variation and the results of selec- 
tion in the uniparental reproduction of D fflugia corona. Genetics, 
1 No. 5. 

Babcock, E. B, and Clausen, R. E. 1918. Genetics in relation to 
agriculture. McGraw-Hill. New York. 

Shamel, A. D., et al. 1918. Citrus fruit improvement. U. S. D. A. 
Buls. 623, 624, 697. 



31 



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